Macrocera analis Lepeletier 1841 Melissodes terminata Smith 1854 Tetralonia bifasciata Smith 1854 Melissodes modesta Smith 1879 Macroglossa oribazi Radoszkowski 1884 Eucera venezuelae Dalla Torre 1896 Melissodes terminata Smith 1854 Thygater chrysophora Holmberg 1903 Macrocera buccosa Vachal 1904 Macrocera buccosa var. nigricollis Vachal 1904 Macroglossapis nigravillosa Crawford 1906 Macroglossapis holmbergi Brèthes 1909 Macroglossapis buccosa var. nigriventris Brèthes 1910 Thygater arechavaletae Bertoni & Schrottky 1910 Thygater zamoranica Cockerell 1949 Thygater ( Thygater ) analis Synopsis of the bee genus Thygater Holmberg 1884 (Hymenoptera, Apidae) in the Brazilian state of Minas Gerais, with the description of a new species and a key to all Brazilian species Freitas, Felipe V. Silveira, Fernando A. Zootaxa 2017 4238 1 1 29 56PPL Lepeletier 1841 Lepeletier 1841 [151,560,1270,1296] Insecta Apidae Thygater Animalia Hymenoptera 5 6 Arthropoda species analis    Macrocera analis Lepeletier 1841;   Melissodes terminata Smith 1854;   Tetralonia bifasciata Smith 1854;   Melissodes modesta Smith 1879;   Macroglossa oribaziRadoszkowski 1884;    Tetralonia pygialisBuysson 1892;    Eucera venezuelaeDalla Torre 1896( nom. nov.for  Melissodes terminata Smith 1854);  Thygater chrysophoraHolmberg 1903;   Macrocera buccosaVachal 1904;   Macrocera buccosavar. nigricollisVachal 1904;  Macroglossapis nigravillosa Crawford 1906;  Macroglossapis holmbergiBrèthes 1909;   Macroglossapis buccosavar. nigriventrisBrèthes 1910;  Thygater arechavaletaeBertoni & Schrottky 1910;  Thygater zamoranicaCockerell 1949;   Thygater( Thygater) analis ; Urban 1967; Urban, Moure & Melo 2012.   Diagnosis.  Thygater analisalso belong to the group of species which females have translucent-amber apical portions of terga and males with entirely black integument and predominantly whitish pilosity on head and mesosoma. In this group, female  T. analiscan be distinguished from those of the sympatric  T. anaeand  T. palliventrisby the presence of bright yellow pilosity on the marginal zones of terga, contrasting with the dark pilosity elsewhere on the terga ( Fig. 5.C) (in  T. anaeand  T. palliventristhe metasomal pilosity is homogeneously yellowish or fuscous ( Fig. 4.B, C, 13.B, C). Mesosomal pilosity of male  T. analismay be predominantly brownish or predominantly whitish; those males with whitish pilosity on mesosoma and predominantly black pilosity on metasoma can be distinguished from male  T. palliventrisonly by the median plate of S7 ( Fig. 3). In  T. analis,the posterior margin of the proximal lobe, and the anterior margin of the distal lobe of S7 are fringed with thick setae ( Fig. 3.A), while in  T. palliventristhese setae are restricted to the proximal lobe ( Fig. 3.D); moreover, the proximal lobe of S 7 in  T. analisis narrow and its anterior and lateral margins meet in a very acute angle ( Fig. 3.A), while in  T. palliventrisit is wider, with its anterior margin bulging and meeting the lateral margin in a wider angle ( Fig. 3.D). The males of  T. analiswith whitish mesosomal pilosity and predominantly black pilosity on metasoma can be distinguished from those of T.  anaeby the absence of a complete marginal fringe on S4 (present in  T. anae) ( Fig. 1).   Description ♀: Integumentblack, except brown on labrum, translucent-amber areas on apex of marginal zones of terga and S2–S5, and dark brown on disk of sterna; on clypeus densely punctate (>1 dp); on mesosoma reticulate with moderately sparse punctation (±1 dp) on mesoscutum, and slightly denser on mesepisternum and scutellum; translucent-amber areas on apex of marginal zones of S2– S5 2/3 as long as F2; on S6 smooth along midlongitudinal stripe. Pilositywhite on gena; on frons and paraocular areas with a mixture of white, fuscous and brown hairs; on vertex fuscous; on mesosoma dense, especially on mesepisternum, its color, especially on dorsum, brown or black or, sometimes, gray; on metasoma, brown basally on T1, black on rest of T1 and basally on T2–T4, depressed, bright yellow on marginal zones of T2–T4 (not evident in some specimens); fuscous and very dense on T5 and T6; on sterna, lighter, varying from fuscous to whitish; on marginal zone of S5, formed by dense hair fringe; on S6 laterally short and dense. Structure—malar area about 1/3 as long as basal mandibular width; basal depression of clypeus reaching 1/6 of clypeal length; clypeus weakly convex, weakly elevated on disc; S6 with weak mid-longitudinal keel.  ♂: Integumententirely black, except whitish-yellow on labrum and translucent-amber on apex of marginal zones of on disc of T1–T5 (not always evidently so) and of S2–S5; on clypeus finely punctate, entirely reticulate; on mesosoma dull, reticulate, punctation moderately dense (1 dp); on disc of T1–T5 sparsely punctate (1 dp); on S6 smooth. Pilosityon head predominantly white with fuscous setae intermingled on paraocular area and frons; on vertex fuscous; on mesosoma and legs variable among specimens, brown, black or rarely gray, on T1 white basally, black apically, as in T2 and T3, and base of T4; on apex of T4, and on T5 and T6 bright yellow (on apex of T4 this golden pilosity may be absent or restricted to a few bristles spread through the postgradular region; in some specimens, this golden pilosity completely absent); dorsally, on meso and metasoma, white, sometimes blackish laterally on sterna; on S6 forming hourglass-shaped, longitudinal band of simple hairs. Structure—malar area 1/2 as long as basal mandibular width; clypeus weakly convex, with weak mid-longitudinal keel, its basal depression occupying 1/5 of clypeal length; T7 deeply emarginated, forming two sharp spines; apical margin of S5 without projections; S6 lacking carinae.   Holotype (male):Brazil (MNHP)—not examined (see taxonomic notes below).   Variation and taxonomic notes:This is the most abundant species in the genus, not only in the state, but throughout South America ( Urban 1967). The striking variation in the pilosity color of  T. analiswas discussed in detail by Urban (1967). In Minas Gerais, both the melanic and the flavinic forms, as defined by Urban, can be found. It is important to note that some of the males have the whole tergal pilosity dark (without the golden hairs found on apical terga of many of them). These males are indistinguishable from male  T. palliventris,except for characters of the terminalia (see identification key and diagnosis, above). The type specimen of  T. analiswas not examined, however, Moure (in Urban 1967) noted that the pilosity of the apical terga of the holotype is pale golden (as opposed to whitish in Lepeletier’s original description), a feature found only in one of the species occurring in Brazil. This gives confidence on the present interpretation of the species.     Examinedmaterial( 227 specimens 101 ♀♀and 126 ♂♂).  ARGENTINA —MISSIONES: Puerto Iguazu—P. N. Iguazu— 04/05/2008, A. Taylor & N. Veiga 1♀; 26/05/20081♀ [PCYU].   BRAZIL— ESPÍRITOSANTO: Dores do Rio Preto— 02/04/2007, A.A. Azevedo 1♀ [UFMG].   GOIÁS: Alto Paraíso de Goiás– 10/04/2007, M.F. Goulart 1♀ [UFMG].   MINAS GERAIS: Alpinópolis—xx/11/1961, C. Elias 1♀[ RPSP]. Alto Capara— 04/04/2007, M.F. Goulart 1♀ [UFMG];   03/12/2015, J.E. Santos-Junior& J.C. Oliveira, 4♀♀ 31♂♂.  Alto Jequitib—PARNA Capara— 03/12/2015, J.E. Santos-Junior & J.C. Oliveira, 7♂♂. Araguari— 16/07/2007, F.A. Silveira 1♀ [UFMG]. Araponga— 25/03/1986, G. Melo 1♂ [UFV];   13/05/1989, A.A. Soares 1♀[UFV].  Belo Horizonte— 02/01/1988, F.A. da Silveira, 9♀♀ 1♂ [UFV]; 25/01/1989, L.A.O. Campos, 3♂♂ [UFV]; 09/03/1996, F.A. Silveira, 3♀♀ 4♂♂ [UFMG]; 19/11/1996, F.A. Silveira 1♂ [UFMG]; 07/01/1997, R.S. Oliveira 1♂ [UFMG]; 08/01/1997, R.M. Carmo 1♂ [UFMG]; 20/01/1997, R.S. Oliveira 1♂ [UFMG]; 08/05/1997, R.S. Oliveira 2♂♂ [UFMG]; 05/10/1997, P.E.M. Guimarães, 3♂♂ [UFMG]; 11/10/1997, P.E.M. Guimarães 1♀ [UFMG]; 19/10/1997, P.E.M. Guimarães 2♂♂ [UFMG]; 25/12/1997, P.E.M. Guimarães, 3♂♂ [UFMG]; 31/12/1997, P.E.M. Guimarães 1♀ [UFMG]; 08/ 08/2000, M. Pompeu 1♂ [UFMG]; 07/06/2000, M. Pompeu 2♀♀ [UFMG]; 21/06/2000, M. Pompeu 1♀ [UFMG]; 26/06/2000, M. Pompeu 1♀ [UFMG]; 22/07/2000, M. Pompeu 1♀ [UFMG]; 23/07/2000, M. Pompeu 1♀ [UFMG]; 08/08/2000, M. Pompeu 1♂ [UFMG]; 09/08/2000, M. Pompeu, 3♂♂ [UFMG]; 21/08/2000, M. Pompeu, 3♂♂ [UFMG]; 30/04/2002, E.A.B. Almeida 1♀ [UFMG]. Brumadinho— 27/12/1997, E.A.B. Almeida 1♂ [UFMG].   Cabo Verde—1920, Diaz 1♂[ MZUSP]. Catas Altas—Serra do Caraça, 11/1961, Kloss, Lenko, Martins & Silva 1♂ [MZUSP]; 20/04/2000, F.A. Silveira 1♂ [UFMG].  Catas Altas da Noruega—  04/01/1989, R.G. Fernandez 1♂[UFV].  Divinópolis— 04/04/1985, E. Morato 1♀ [UFV];   07/04/1985, G. Melo 1♀[UFV].  Gonçalves— 26/03/2007, F.A. Silveira 1♀ [UFMG]; 27/03/2007, F.A. Silveira, 4♂♂ [UFMG]. Governador Valadares— 27/06/1985, G. Melo 1♂ [UFV]; 15/02/1986, G. Melo 1♂ [UFV];   16/02/1986 1♀ 2♂♂[UFV].  Grão Mogol— 22/01/2005, A.A. Azevedo 1♀ 1♂ [UFMG]. Guiricema— 05/08/2011, R.V. Silva 1♀ [UFMG]. Igarap— 23/02/1086, G. Melo 2♀♀ 2♂♂ [UFV]; 20/11/2006, C.L. Yurrita 1♀ [UFMG]. Ipuiuna— 21/02/2012, L. Patoca 1♀ [RPSP].  Itabirito—  01/01/1987, F.A. Silveira 1♂[UFV].  Itamarandiba— 17/01/1998, F.A. Silveira 1♀ [UFMG]. Itamonte— 20/10/2006, A.A. Azevedo 2♂♂ [UFMG]; 20/10/2006, M.F. Goulart 1♀ 1♂ [UFMG]. Itaúna— 27/12/1997, P.E.M. Guimarães 1♀ [UFMG]. Joaquim Felício— 31/03/2005, A.A. Azevedo & F.A. Silveira 1♀; 20/05/2006, M.F. Goulart 1♀ [UFMG].   Juiz de Fora—  11/03/1985, M. Garcia 1♂[UFV].  Moeda— 27/01/1996, F.A. Silveira 1♀ [UFMG].   Morro do Pilar—  08/10/2014, E.L.N. Siqueira 1♂[ UFMG]. Nova Lima— 20/01/1998, E.A.B. Almeida 1♂ [UFMG]; 13/11/2004, N.G. Fonseca 1♀ [UFMG]; 18/11/2004, N.G. Fonseca 1♀ [UFMG]. Paraopeba—Floresta Nacional de Paraopeba, 02/12/1986, F.A. Silveira 1♀ [UFV]; 02/12/ 1986, F.A. Silveira 1♂ [UFMG]; 03/08/1987, F.A. Silveira 1♀ [UFV]; 31/08/1987, F.A. Silveira 1♀ [UFMG]; 11/ 10/1987, F.A. Silveira 1♀ [UFV]; 21/10/1998, V. Silva 1♂ [UFMG]. Passos—04/1964, C. T. Elias 1♀ 4♂♂ [MZUSP]. Patrocínio— 08/12/2007, J.E. Santos-Junior 1♀ [UFMG].  Ponte Nova—BR–120, km 493  14/02/1989, F.A. Silveira 1♂[UFV]; Fazenda EPAMIG  10/11/1989, F.A. Silveira 2♂♂[UFV].  Ritápolis— 19/01/1974, J.F. Camargo 1♀ [RPSP]. Sabar— 07/04/1996, F.A. Silveira 2♀♀ [UFMG]; 28/11/1999, F.A. Silveira 1♀ [UFMG]. Santana do Riacho— 09/01/2005, A.A. Azevedo 1♀ [UFMG].   São Gonçalo do Rio Abaixo—  01/01/2000, E.A.B. Almeida 1♂[ UFMG];   17/01/2004, L.R.R. Faria Jr. 1♂[ UFMG].  São Gonçalo do Rio Preto— 13/09/2003, Y. Antonini 1♀ [UFMG]. São Roque de Minas— 24/02/2001, E.A.B. Almeida 1♀ [UFMG]. Taiobeiras— 02/01/ 1998, S.T.M. Guerra 1♂ [UFMG]. Urucuia— 16/11/2008, J.F. Camargo 1♀ [RPSP]. Viçosa— 15/04/1928, P.H. Rolfs 1♀ [UFV];   20/05/1932, E.J. Hambleton 1♀[UFV]; CampusUFV,   06/01/1988, J.R. Cure 2♂♂[UFV]; Campus UFV  10/11/1988, M.J. Fachini 2♂♂[UFV]; CampusUFV, 03/12/1988, G.S. Bastos Fo. 1♀ [UFV]; Campus UFV, 08/12/1988, G.S. Bastos Fo.  1♂[UFV]; CampusUFV,   08/12/1988, J.R. Cure 2♂♂[UFV]; Campus UFV  13/12/1988, M.J. Fachini 1♀[UFV]; Campus UFV  18/01/1989, M.J. Fachini 1♀[UFV]; CampusUFV  11/ 02/1989, G.S. Bastos Fo. 2♀♀ [UFV]; Campus UFV 19/02/1989G.S. Bastos Fo.  1♂[UFV]; Campus UFV  24/02/ 1989, M.J. Fachini 1♂[UFV]; CampusUFV,   04/03/1989, G.S. Bastos Fo. 1♀[UFV]; Campus[UFV],   04/03/1989, M.J. Fachini 1♀ 1♂[UFV]; CampusUFV  11/03/1989 G.S. Bastos Fo.  1♂[UFV]; Campus UFV  11/03/1989, J.R. Cure 1♂[UFV]; Campus UFV  11/03/1989, M.J. Fachini 2♀♀[UFV]; CampusUFV  18/03/1989, G.S. Bastos Fo. 1♀ [UFV]; 08/11/1989, M. Thiengo 1♂ [UFV]; 27/11/2001, M.S.O. Leite 1♂ [UFMG]; 30/11/2001, M.S.O. Leite 1♀ [UFMG]; 04/12/2001, M.S.O. Leite 1♂ [UFMG];   05/12/2001, M.S.O. Leite 1♀[ UFMG].  PARAÍBA: Remigio— 18/06/2011, S.C. Aguiar 1♂ [UFMG]; 15/07/2011, A.S. Silva 1♀ 1♂ [UFMG];   02/10/2011, S.C. Aguiar 1♀[ UFMG].  PERNAMBUCO:  Cabo de Santo Agostinho— 28/09/2001, C. Schlindwein 1♂ [UFMG].   Camaragibe—  09/04/2000, C. Schlindwein 1♂[ UFMG].  RIO DE JANEIRO: Nova Friburgo—   19/03/2012, T.M.S. Mesquita 1♀[ UFMG];  Rio de Janeiro—   28/09/2013, P.C. Zama, 4♀♀[ UFMG].  RIO GRANDE DO SUL: Candiota— Chácara das Hortec.—   23/02/2000, S. Witter 1♂[ PUC–RS];  Esmeralda— 05/01/2011, R. Halinski 1♂ [PUC–RS]; Guarani das Missões— 23/07/2012, F. Tirelli 1♀ 1♂ [PUC–RS]; 30/06/2012, S. Witter 1♀ [PUC– RS];   31/06/2012, S. Witter 1♂[ PUC–RS].  Osório— 21/03/1993, I. Alves dos Santos 1♂ [PUC–RS].   SANTA CATARINA: São Bonifácio— 17/11/2015, F.V. Freitas 1♂ [UFMG]. São Francisco do Sul— 20/12/2011, C.F. Cardoso, 4♀♀ 4♂♂ [UFMG].   SÃO PAULO: Campinas— 07/06/1987, R.B. Martins 1♂ [UFMG].   Distribution in Minas Gerais:  Thygater analisis probably found all over the state. Besides the localities cited above for the examined specimens, the following records were given by Urban (1967): Arax, Cássia, Cataguases, Ibi, Ibiraci, Jacu, Nova Resende, Poços de Caldas, Rio Piracicaba, Tapira, Timóteo, Varginha.  Flower records: Alismataceae—  Echinodorus grandiflorus,Sertãozinho–SP [RPSP];  Bignoniaceae—  Podranea ricasoliana,BeloHorizonte—MG [UFMG];  Clytostomasp., Belo Horizonte—MG [UFMG];  Tecoma stans,Belo Horizonte–MG [UFMG];  Bixaceae—  Bixa orellana,Sabar–MG [UFMG].  Brassicaceae—  Brassica napus, Guaranidas Missões—RS [PUC–RS]. Curcubitaceae—  Cucurbita pepo, Igarap—MG [UFMG].  Fabaceae—unidentified species, Taiobeiras—MG [UFMG].  Lamiaceae—  Hyptidendron canum,BeloHorizonte—MG [UFMG];  Salvia brevipes, Candiota—RS [PUC–RS].  Lythraceae—  Cupheasp., Catas Altas – MG [UFMG].  Melastomataceae—  Trembleya phlogiformisDC Central Amazon [Renner 1984 apud Renner 1989].  Myrtaceae—  Psidium guajava,BeloHorizonte—MG [UFMG]. Myrcia racemosa, Ubatuba—SP [ Fidalgo & Kleinert 2009].  Ochnaceae—  Ouratea casteneifolia,Belo Horizonte[UFMG].  Passifloraceae—  Passiflorasp., Sabar—MG [UFMG].  Solanaceae—  Solanum cernuum,Viçosa[Cure et al.1993];  Solanum lycocarpum,Paraopeba[ Silveira 1989];  Solanumsp., Paraopeba [ Silveira 1989].  Verbenaceae—  Duranta repens,BeloHorizonte—MG [UFMG]; Stachytarphettasp., Moeda –MG [UFMG]. 1437764180 [199,1011,404,429] Examined Argentina ARGENTINA 9 10 454 101 126 1437764166 Brazil Rio Preto 9 10 1 ESPIRITO SANTO 1437764159 [766,1124,476,501] Brazil GOIAS 9 10 1 GOIAS 1437764171 [311,1049,511,536] RPSP Alpinopolis & C. Elias Brazil MINAS GERAIS 9 10 1 1 MINAS GERAIS 1437764156 [418,1316,548,573] 2015-12-03 J. E. Santos-Junior & J. C. Oliveira Brazil Alto Jequitib 9 10 35 4 31 MINAS GERAIS 1437764172 [673,1274,620,645] 1989-05-13 A. A. Soares Brazil Belo Horizonte 9 10 1 1 MINAS GERAIS 1437764176 [264,718,980,1005] MZUSP Diaz Cape Verde Cabo Verde 9 10 1 1 1437764177 1989-01-04 R. G. Fernandez Norway Divinopolis 9 10 1 1 1437764178 1985-04-07 G. Melo Norway Goncalves 9 10 1 1 1437764174 1986-02-16 Norway Grao Mogol 9 10 3 1 2 1437764157 [306,1059,1231,1256] 1987-01-01 F. A. Silveira Norway Itamarandiba 9 10 1 1 1437764162 [750,1407,1340,1365] 1985-03-11 Juiz de Fora & M. Garcia Norway Moeda 9 10 1 1 1437764158 [633,1356,1375,1400] 2014-10-08 UFMG E. L. N. Siqueira Norway Morro do Pilar 9 10 1 1 1437764161 1989-02-14 1989-11-10 1989-02-14 F. A. Silveira Norway Ritapolis 9 10 3 3 1437764164 2000-01-01 UFMG Sao Goncalo do Rio Abaixo & E. A. B. Almeida Norway Rio Abaixo 9 10 1 1 1437764179 [421,1239,1700,1725] 2004-01-17 UFMG L. R. R. Faria Jr. Norway Rio Preto Sao Goncalo do Rio Preto 9 10 1 1 1437764167 [365,1002,1807,1832] 1932-05-20 E. J. Hambleton Norway Campus 9 10 1 1 1437764173 1988-01-06 1988-11-10 1988-01-06 J. R. Cure & M. J. Fachini Norway Campus 9 10 4 4 1437764169 [639,926,1880,1904] Norway Campus 9 10 1 1 1437764168 1988-12-08 1989-02-11 1988-12-08 J. R. Cure & M. J. Fachini & G. S. Bastos Fo Norway Campus 9 10 4 2 2 1437764175 1989-02-24 M. J. Fachini Norway Campus 9 10 2 2 1437764170 [664,1290,1988,2012] 1989-03-04 G. S. Bastos Fo. Norway Campus 9 10 1 1 1437764165 1989-03-04 1989-03-11 1989-03-04 M. J. Fachini & G. S. Bastos Fo Norway 10 11 Campus 9 10 2 1 1 1437764163 1989-03-11 1989-03-18 1989-03-11 J. R. Cure & M. J. Fachini & G. S. Bastos Fo Norway Campus 10 11 4 2 2 1437764160 2001-12-05 UFMG M. S. O. Leite Norway Remigio 10 11 1 1 PARAIBA 1437764203 2011-10-02 UFMG S. C. Aguiar & Cabo de Santo Agostinho Norway Cabo de Santo Agostinho 10 11 1 1 PERNAMBUCO 1437764204 [151,1190,367,392] 2000-04-09 UFMG Camaragibe & C. Schlindwein Norway Nova Friburgo 10 11 1 1 RIO DE JANEIRO 1437764188 2012-03-19 UFMG T. M. S. Mesquita Norway Rio de Janeiro 10 11 1 1 Rio de Janeiro 1437764182 2013-09-28 UFMG P. C. Zama & Chacara das Hortec. Norway Candiota 10 11 4 4 RIO GRANDE DO SUL 1437764197 [544,1098,439,464] 2000-02-23 PUC S. Witter Norway Esmeralda 10 11 1 1 RIO GRANDE DO SUL 1437764198 [215,749,511,536] 2012-06-31 PUC S. Witter Norway Osorio 10 11 1 1 RIO GRANDE DO SUL 1437764181 Norway SANTA CATARINA 10 11 1 SANTA CATARINA 1437764187 [498,773,583,608] Norway SAO PAULO 10 11 1 SAO PAULO